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Genetic Diversity and Genetic Heterogeneity of Bigfin Reef Squid “Sepioteuthis Lessoniana” Species Complex in Northwestern Pacific Ocean

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Genetic Diversity and Genetic Heterogeneity of Bigfin Reef Squid “Sepioteuthis Lessoniana” Species Complex in Northwestern Pacific Ocean 8 Genetic Diversity and Genetic Heterogeneity of Bigfin Reef Squid “Sepioteuthis lessoniana” Species Complex in Northwestern Pacific Ocean Hideyuki Imai and Misuzu Aoki University of the Ryukyus Nara Women’s University Japan 1. Introduction The bigfin reef squid Sepioteuthis lessoniana Férussac, 1831 in Lesson (1830–1831) is widely distributed in the Indo-Pacific, where it is a very valuable fishery resource (Dunning, 1998). Thus, a lot of ecological research of this species were reported (e.g. Ikeda, 1933; Choe & Ohshima, 1961; Segawa, 1987; Ueta, 2003; Ikeda et al., 2009). Segawa et al. (1993a; 1993b) showed that within Sepioteuthis lessoninana have diferrences of egg chracteristics and reproductive trait in Ishigakijima Island. Izuka et al. (1994) reported an allozyme analysis found so-called S. lessoniana around Ishigakijima in Okinawa Prefecture, Japan, includes at least three biological species (Figure 1 & 2). Local fishers call the three species “aka-ika,” which has a red body, “shiro-ika” or “aori-ika,” which has a white body, and “kua-ika,” which is smaller than the other two. Of these, the range of “shiro-ika” extends to the coast of the main Japanese islands. This is the extent of its taxonomic classification thus far. This is due in part to the limited number of distinguishing morphological characters but also because the type specimens is no longer available and type locality has not been disignated (Lu et al., 1995; Jereb & Roper, 2006). This makes it difficult to determain whether genetically recognized species are undescraibed species or one of 13 known synonymies (Young, 2002). In this study, we treated “aka-ika” as Sepioteuthis sp. 1, “shiro-ika” as Sepioteuthis sp. 2, and “kua-ika” as Sepioteuthis sp. 3. A previous population genetics study found significant differences in the genetic heterogeneity of Sepioteuthis sp. 2 between Pacific Ocean and Japan Sea populations using allozyme analysis (Yokogawa & Ueta, 2000). Yokogawa and Ueta (2000) did not include the Okinawan Sepioteuthis sp. 2 population in their study. In addition, Pratoomchat et al. (2001) found no significant genetic heterogeneity between Japanese and Thai Sepioteuthis sp. 2 populations, while our present study tried significant differences in the genetic heterogeneity of the Japanese and Vietnumese Sepioteuthis sp. 2 populations. Recently, Aoki et al. (2008a) reported significant genetic heterogeneity between Japanese and Vietnumese populations of Sepioteuthis sp. 2 using DNA sequencing analysis of the mitochondrial noncoding region. Therefore, this study examined the genetic diversity (i.e., the average heterogeneity) and gene flow among Sepioteuthis sp. 2 populations using allozyme analysis and among www.intechopen.com 152 Analysis of Genetic Variation in Animals populations of Sepioteuthis sp. 1 and Sepioteuthis sp. 3 using mitochondrial DNA noncoding region sequencing of populations from Japanese, Taiwanese, and Vietnamese waters. A B C D E F Fig. 1. A: Sepioteuthis sp. 1, B: Sepioteuthis sp. 2, C: Sepioteuhis sp. 3, D: Sepioteuthis sp. 1 egg capsules with 5-13 (mean = 9) per capsule, E: Sepioteuthis sp. 2 egg capsules with 3-8 eggs (mode = 6) per capsule and F: Sepioteuthis sp. 3 egg capsules with consistently two eggs per capsule laid under dead table coral in shallow waters. Black bar indicated 50mm in length. 2. Materials & methods 2.1 Allozyme analysis of Sepioteuthis sp. 2 We collected 327 adults between September 1998 and June 2006 from Noto, Ishikawa, Japan (83 individuals), Mugi, Tokushima, Japan (51), the Goto Islands, Nagasaki, Japan (58), Nakagusuku, Okinawa (52), Keelung, Taiwan (23), and the Gulf of Tonkin, Vietnam (60). All www.intechopen.com Genetic Diversity and Genetic Heterogeneity of Bigfin Reef Squid “Sepioteuthis lessoniana” Species Complex in Northwestern Pacific Ocean 153 Fig. 2. Electrophoretic patterns of asparate aminotransferase (AAT) of Sepioteuthis lessoniana complex. Lane 1-2: Sepiteuthis sp. 2, Lane 3-4: Sepioteuthis sp. 3 and Lane 5: Sepioteuthis sp. 1. These three species are clearly identified by the Aat-1* marker (Izuka et al. 1994). specimens were fresh and immediately sent to a refrigerator in the laboratory. The buccal bulb muscle was removed and kept frozen at –40°C until the allozyme analysis. Small pieces of liver and skeletal muscle were dissected from selected specimens and minced individually in an equal volume of distilled water on ice. Electrophoresis was conducted in a glass box with ice on top of it. The box was in a refrigerator at a constant voltage (250 V) until the Amido Black 10B marker moved seven cm from the origin. The allozymes were tested using 12.5% horizontal starch–gel electrophoresis and the two buffer systems described by Clayton and Tretiak (1972) and modified by Numachi (1989): citric acid N-(3-aminopropyl) diethanolamine (CAEA, pH 7) and citric acid N-(3-aminopropyl) morpholine (CAPM, pH 6). Each gel was sliced into six 1-mm-thick sheets with a wire gel cutter (Numachi, 1981) and stained for the enzymes aspartate aminotransferase (AAT), isocitrate dehydrogenase (IDHP), lactate dehydrogenase (LDH), phosphoglucomutase www.intechopen.com 154 Analysis of Genetic Variation in Animals (PGM), and phosphogluconate dehydrogenase (6PGD) according to Shaw and Prasad (1970), Numachi (1970a, b), and Taniguchi and Numachi (1978). The locus and gene nomenclature followed Shaklee et al. (1990). Polymorphisms involving several alleles with frequencies of more than 5% were tested at a significance level of 0.05 to determine whether they were consistent with Hardy–Weinberg equilibrium. The average heterozygosity H (Nei, 1978) was calculated as a measure of genetic diversity. The χ2 homogeneity test of allele frequency among samples was also performed. 2.2 Mitochondrial non-coding region of Sepioteuthis sp. 1 and sp. 3 In total, 116 Sepioteuthis sp. 1 were collected between April 2005 and September 2006 at three localities: Itoman, Okinawajima (49 individuals), Ishigakijima (38), and Keeling, Taiwan (29). An arm or part of the mantle muscles was kept in 90% ethanol and DNA was extracted with TNES 8M-Urea buffer. For Sepioteuthis sp. 3, 60 samples were collected between October 2005 and July 2006 from Nago, Okinawajima (30), and Ishigakijima (30). Crude DNA was extracted by TNES 8M Urea buffer and proteinase K digestion followed by a phenol- chloroform isoamyl method described Imai et al. (2004). We analyzed the noncoding region 2 (NC2) between the Ala and Trp transfer RNAs (tRNAs). The original primers SL-Ala (5'-GGTAACCCTTTCTGTATGATTGC-3') and SL-Trp (5'-AAAGACCTTGAAAGTCTTCAG-3'), which target a portion of tRNA-Ala and tRNA- Trp, respectively, were used with the polymerase chain reaction (PCR) to amplify NC2 (Aoki et al., 2008a). The PCR reactions were performed using BIOTAQ (Bioline, UK). A GeneAmp 9700 (Applied Biosystems, USA) thermal cycler was used with the following setting: 94°C for 120 s, followed by 30 cycles at 94°C for 30 s, 60°C for 30 s, and 72°C for 45 s. The PCR products were purified using a PCR Product Pre-sequencing Kit (USB, USA). The nucleotide sequences were determined using ABI 3700 (Applied Biosystems, USA) genetic analyzers. All sequences were initially aligned using ClustalX ver. 1.83.1 (Thompson et al., 1997) and then edited manually using MacClade4 ver. 4.08 (Maddison and Maddison, 2005). The haplotype diversity h (Nei, 1987) and nucleotide diversity π (Tajima, 1983) within populations were calculated using Arlequin ver. 2.000 (Schneider et al., 2000). An analysis of molecular variance (AMOVA; Excoffier et al., 1992) was used to test the population structure within species for Sepioteuthis sp. 1 using Arlequin. For Sepioteuthis sp. 3, AMOVA could not be performed because there were fewer than three localities. Therefore, homogeneity was tested using the chi-square randomization method (Monte Carlo simulation) with 100,000 randomizations of the data (Roff and Benzen, 1989). Significance thresholds were Bonferroni-corrected for multiple pairwise comparisons. Relationships of haplotypes were assessed using a minimum spanning tree created via the Minspanet algorithm in Arlequin and drawn by hand. 3. Results and discussion 3.1 Allozyme analysis of Sepioteuthis sp. 2 Regarding the eight loci for the six enzymes analyzed, the five loci Aat-1*, Idhp-1*, Ldh-1*, Mdh-1*, and Mdh-3* showed no differences among and within localities, and no genetic www.intechopen.com Genetic Diversity and Genetic Heterogeneity of Bigfin Reef Squid “Sepioteuthis lessoniana” Species Complex in Northwestern Pacific Ocean 155 polymorphism was recognized. Two Mdh-2* heterozygotes were found in Vietnam, although the frequency was 0.017; therefore, it was not considered a polymorphic allozyme locus (Table 1). Genetic polymorphism was detected within a locality for Pgm* and 6pgd*. The polymorphic allozyme loci were in Hardy–Weinberg equilibrium at the localities. Most of the alleles linked to a locus were monomorphic. A marked excess of homogeneity was found. The average observed heterozygosity H=0.005–0.052 was similar to the values of H=0.037 reported by Izuka et al. (1996) and 0.052–0.070 by Yokogawa and Ueta (2000). Other loliginid species have similar heterozygosity values: Loligo pealeii, H=0.006; Lolliguncula brevis, H=0; L. plei, H=0 (Garthwaite et al., 1989); Ommastrephes bartrami, H=0.004; Sthenoteuthis oualaniensis, H=0.011; Todarodes pacificus, H=0.043; Loliolus japonica, H=0.030 (Fujio & Kawada, 1989); L. vulgaris reynaudii, H=0.030; L. gahi, H=0.059 (Carvalho & Loney, 1989); L. bleekeri, H=0.003 (Suzuki et al., 1993); and L. chinensis, H=0.006–0.009 (Yeatman and Benzie, 1993). The family Loliginidae appears to be characterized by low genetic diversity. Okinawajima Table 1. Allele frequencies at eight loci and indices of genetic heterozygosities within six localities of Sepioteuthis sp. 2. No allele frequency gap was observed among different localities for the polymorphic allozyme loci 6pgd* allele frequency. In contrast, a significant difference was detected between Pgm* in the Japanese and Vietnamese localities (Table 2).
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